On humans and wildlife in Mediterranean islands

Aim To investigate the effects of human‐induced landscape changes in Mediterranean islands on the ecological and evolutionary responses of bird communities and populations. The combination of mass extinction of large mammals and massive deforestation by humans was hypothesized to produce new selection regimes to which organisms were likely to respond. Habitat selection and niche breadth have been investigated at the scale of species, and phenotypic variation at the scale of local populations. Location The study was carried out along habitat gradients and in habitat mosaics at different spatial scales on the island of Corsica and in areas of similar size and structure in continental France. Methods Two sets of gradients have been used for investigating habitat selection and niche breadth: gradients of altitude, and gradients of vegetation structure. Population studies focused on the blue tit, Cyanistes caeruleus. Large samples of breeding attempts by this species in 10 habitats provided detailed data on phenotypic variation of fitness‐related traits both on Corsica and on the mainland. Results The extent of niche space used by birds differed substantially depending on which habitat gradient was considered. Many species have been found to contract their habitat niche along the elevation gradient on Corsica compared with the mainland, whereas all species in the vegetation gradient broadened their niche on the island. Breeding patterns of the blue tit differed considerably depending on whether they settle in deciduous oaks (Quercus humilis) or in evergreen sclerophyllous oaks (Quercus ilex). Phenotypic variation of breeding traits was much higher on the island, where more populations were correctly timed for the best breeding period than on the mainland, a pattern that is likely to result from lower dispersal of organisms on the island. Main conclusions The differences in observed niche breadth between the two series of habitat gradients is explained both by the species‐specific ecology of the species and the human‐induced environmental history of Corsica. Large‐scale landscape changes provided new opportunities for island colonization by non‐forest species, which are isolated as small, ‘fugitive’ local populations. In both gradients, forest species that are typical components of the Corsican bird fauna definitely expanded their niche and occupied a wider range of habitats on Corsica than on the mainland. At the population scale, landscapes included habitat patches with contrasted selection regimes, which resulted in high phenotypic variation for many fitness‐related traits. Reduced dispersal of birds on the island resulted in a much higher degree of local differentiation on Corsica than on the mainland.     

Patterns of Genetic and Reproductive Traits Differentiation in Mainland vs. Corsican Populations of Bumblebees

Populations on islands often exhibit lower levels of genetic variation and ecomorphological divergence compared to their mainland relatives. While phenotypic differentiation in characters, such as size or shape among insular organisms, has been well studied, insular differentiation in quantitative reproductive traits involved in chemical communication has received very little attention to date. Here, we investigated the impact of insularity on two syntopic bumblebee species pairs: one including species that are phylogenetically related (Bombus terrestris and B. lucorum), and the other including species that interact ecologically (B. terrestris and its specific nest inquiline B. vestalis). For each bumblebee species, we characterized the patterns of variation and differentiation of insular (Corsican) vs. mainland (European) populations (i) with four genes (nuclear and mitochondrial, 3781 bp) and (ii) in the chemical composition of male marking secretions (MMS), a key trait for mate attraction in bumblebees, by gas chromatography-mass spectrometry (GC-MS). Our results provide evidence for genetic differentiation in Corsican bumblebees and show that, contrary to theoretical expectations, island populations of bumblebees exhibit levels of genetic variation similar to the mainland populations. Likewise, our comparative chemical analyses of MMS indicate that Corsican populations of bumblebees are significantly differentiated from the mainland yet they hold comparative levels of within-population MMS variability compared to the mainland. Therefore, insularity has led Corsican populations to diverge both genetically and chemically from their mainland relatives, presumably through genetic drift, but without a decrease of genetic diversity in island populations. We hypothesize that MMS divergence in Corsican bumblebees was driven by a persistent lack of gene flow with mainland populations and reinforced by the preference of Corsican females for sympatric (Corsican) MMS. The impoverished Corsican bumblebee fauna has not led to relaxation of stabilizing selection on MMS but to consistent differentiation chemical reproductive traits on the island.     

Macrogeographic variations in food choice of mainland citril finches Carduelis [citrinella] citrinella versus insular Corsican (citril) finches Carduelis [citrinella] corsicanus

We studied variations in diet composition in sub-populations of mainland citril finches Carduelis [citrinella] citrinella in European mountain chains, compared to closely related insular Corsican (citril) finches Carduelis [citrinella] corsicanus on several Mediterranean Islands with a special emphasis on key food plants used during the breeding period. We found that citril finch sub-populations of the Pre-Pyrenees and the Black Forest rely mostly on mountain pine and dandelion seeds as main food resources during the breeding season. In contrast, insular Corsican finches on the Mediterranean islands of Capraia and Corsica fed mainly on various herb species. Finches in Corsica fed predominantly on shepherd’s purse, while birds from Capraia were choosing rosemary as their main food plant during the breeding period. These observations confirmed the previously expected difference in food choice between the two forms due to the observed increased niche breadth on Corsica. However, Corsican finches on Sardinia showed a food choice more similar to that of the citril finch populations, in that they fed predominantly on black pine seeds during breeding season. Thus we were not able to find clear behavioural differences in diet composition between all studied sub-populations of citril and Corsican finches. To conclude, food choice of both forms is highly adaptive and, in the different breeding areas, distinct key plants may be of special importance during the breeding season. The preference for these plants is likely to be linked to their abundance and their energetic and nutritional content.     

Conservation biogeography of large Mediterranean islands. Butterfly impoverishment, conservation priorities and inferences for an ecologica "island paradigm"

Conservation biogeography is considered the Cinderella of biological conservation. Nevertheless biogeography provides the basis for establishing species distributions over space and time, therefore conservation priorities among areas and individual species. We demonstrate that there is no need to simplify analyses by using subsets of species (rare species, endemics) as surrogates. In doing so, we apply strict biogeographical techniques to determine butterfly impoverishment on three of the west Mediterranean's largest islands (Sardinia, Corsica and Sicily). The analyses performed on species, both collectively and individually, reveal that regional species richness in the Mediterranean zone can be largely predicted by latitude, altitude and latitudinal range (maximum minus minimum latitude), but that Sardinia and Corsica have clearly impoverished faunas. Logistic regression at individual species level demonstrates that several species, predicted to be present in these islands on the basis of their continental distributions, are actually absent. When compared with species that are present in these islands, such missing species are disclosed as having ecological traits which reduce their colonization capability. Probabilities of occurrence are calculated for each species on each island; they reflect the potential for each butterfly species to migrate to and colonise each island, and can be considered as a measure of conservation value. As such, species present on islands but having low immigration probabilities are predicted to represent isolated populations from the mainland that are unlikely to re-colonize the islands in the case of extinction. Island endemic species and races are shown to have lower occurrence probabilities compared to widespread species occurring on islands and illustrate the usefulness of occurrence probabilities for identifying isolated populations in need of conservation attention.     

Bird assemblages on Amazonian river islands: Patterns of species diversity and composition

The principles of island biogeography are rarely applied to the animal assemblages of Amazonian river islands. Here, we compare bird assemblages of Amazonian river islands with a variety of mainland habitats. We also examine how bird species diversity and composition are related to island physical attributes. Birds were sampled with mist nets and qualitative censuses on 11 river islands and 24 mainland sites on the lower reaches of the Rio Negro in the Brazilian Amazon. Island bird assemblages were characterized by lower species richness and a higher abundance of a few dominant species. Additionally, the species composition of the islands was distinct from that of the mainland, including the nearby floodplain habitats. The number of bird species increased with island size and habitat diversity, and decreased with degree of isolation. In addition, small islands tended to harbor an impoverished subset of the species present on larger ones. Bird species diversity and composition on Amazonian river islands are likely influenced by the ecological succession and historical events affecting island formation. Considering their small total area across the Amazon basin, these insular fluvial communities could be disproportionately threatened by river channel disturbances related to climate change or hydroelectric dam development. Abstract in Portughese is available with online material.     

Why do Corsican Blue Tits Cyanistes caeruleus ogliastrae not use nest boxes for roosting?

We compared winter roosting behaviour of Blue Tits Cyanistes caeruleus in the mainland of southern France and on the island of Corsica in small-holed nest boxes. While in southern France Blue Tits use nest boxes for roosting, Corsican Blue Tits do not. We suggest that this behaviour is innate because Corsican wild-caught as well as F1 and F2 birds born in captivity do not use nest boxes in aviaries even when kept adjacent to mainland tits that sleep in boxes. We suggest that the cost/benefit balance differs between mainland France and Corsica: in Corsica, the costs of roosting in the canopy are less than in mainland France because (1) predators, e.g. owls, are rare in Corsica while abundant on the mainland, (2) the permanent evergreen foliage of the dominant tree in Corsica, the holm oak, could be preferred to nest boxes which are not risk-free, and (3) the costs of using cavities for roosting could be higher in Corsica because of increased mammal predator pressure and higher ectoparasite load than on the mainland.     

Edaphic beetle (Insecta: Coleoptera) diversity over a forest-matrix gradient in a tropical rainforest

Species from natural communities show different capacities for moving across contrasting habitats, and they may gradually respond along the transition between forests and anthropogenic areas. Because beetles are effective bioindicators, we analyzed the Coleoptera assemblage structure in the transition between sugarcane matrices and forest fragment. The study was conducted in an Atlantic rainforest fragment and sugarcane matrices after 2 and 6 months of replantation. Beetles were sampled in linear transects that were 50, 100, and 200 m from the forest edge and toward both forest core and matrices. We analyzed beetle abundance, species richness and composition, and diversity numbers. The abundance and species richness were higher in the forest, and species richness was higher at the sampling site 100 m inside the forest than at the gradients within the sugarcane matrices. The species composition differed between the forest and matrices, but not between the matrices with different replantation ages. Alpha diversity based on the number of species was higher in the forest, and alpha diversity based on the Shannon index was higher in the forest and matrix after 6 months of replantation. Our results suggest that the sugarcane matrices, chiefly the matrix after 2 months of replantation, sustain an impoverished beetle assemblage when compared with the forest fragment. Despite the strong habitat distinction, the beetle fauna in the sugarcane matrices was not affected by the horizontal distance from the forest. Here, we have highlighted the importance of forest fragments embedded in harsh matrices for the maintenance of biodiversity.     

Skipper impoverishment on large West Mediterranean islands (Lepidoptera Hesperioidea): deterministic,historical and stochastic factors

Biogeographical analyses are applied to skipper (Hesperioidea) presence/absence data from the Western Mediterranean mainland and the three largest islands (Sardinia, Corsica and Sicily) in order to identify potential conservation issues. The analyses performed on species, both collectively and individually, indicate that regional species richness and occurrence in the Mediterranean zone are largely predicted by latitude and area but that islands have impoverished faunas. Several species, predicted to be present on these islands from logistic regression of their continental distributions, are actually absent. The number of species predicted to be present from logistic regression analyses for each island, closely matched the number of species predicted to occur in regional-focused multiple regression analysis. This suggests that missing species have been identified. When compared with species that occur in Sicily and Corsica, the missing species are shown to differ for ecological traits, mainly those linked to altitudinal tolerance. No ecological distinctions were disclosed for Sardinian skippers suggesting a mainly stochastic colonisation. These results, and those from an analogous study carried out on Papilionoidea, point to Hesperioidea having (i) overall more impoverished faunas on islands and (ii) being subject to stochastic or historical colonisation events more than Papilionoidea. Species not predicted to occur on islands based on their mainland distributions and ecological traits, are foci for conservation attention. However, as many species becoming extinct on the islands may be irreplaceable, all species, in particular the Sardinian ones, deserve to be conserved.     

Long-Term Trends in Faunal Recolonization After Bauxite Mining in the Jarrah Forest of Southwestern Australia

Abstract Fauna serve a key role in many forest ecological processes. Despite this, few studies have considered long‐term faunal recolonization after mining and rehabilitation of forest ecosystems. In the jarrah forest of southwestern Australia, permanent fauna monitoring sites have been established in bauxite mined areas rehabilitated in 1990 and in a range of representative unmined forest control sites. At each site mammals, birds, reptiles, and ants were surveyed in 1992, 1995, and 1998. The aims of the monitoring were to develop a better understanding of faunal recolonization trends, to produce recommendations for promoting fauna return, and to consider which techniques and fauna groups are best suited for monitoring recolonization. The results showed that successional trends varied between fauna groups. Generalist foraging mammals recolonized rapidly, whereas small predators took longer. Feral mice were initially abundant and then declined. Birds gradually recolonized, and after 8 years bird communities were very similar to those in unmined forest sites. Reptile species took longer, and after 8 years numbers of species remained lower than in unmined forests. Species richness and diversity of ants in 8‐year‐old rehabilitation were comparable with those of unmined forest in some rehabilitated sites but were lower in others. The composition of ant communities was still different from that of unmined sites. Ant species that only use disturbed forest declined rapidly in abundance as rehabilitation aged. The results suggest that although the rates of faunal recolonization will vary, with time most or all mammal, bird, reptile, and ant species should inhabit rehabilitated bauxite mines. The densities of many are likely to be similar to those in unmined forest, but for others it is too early to know whether this will be the case. Techniques for promoting fauna return are discussed. This study demonstrates that no single fauna group is suitable for use as an overall “indicator” of faunal recolonization; different fauna species and groups reflect different aspects of faunal succession.     

Assemblage-level responses of phyllostomid bats to tropical forest fragmentation: land-bridge islands as a model system

Aim Working within a system of high structural contrast between fragments and the surrounding matrix, we assessed patterns of species loss and changes in species composition of phyllostomid bats on artificial land‐bridge islands relative to mainland assemblages, and evaluated the responses of bats to forest edges. We further examined the relative influence of local‐scale characteristics (e.g. vegetation structure, island area) versus landscape attributes (e.g. forest cover, patch density) and the importance of spatial scale in determining phyllostomid species richness and composition on islands. Location Islands in Gatún Lake and adjacent mainland peninsulas in the Barro Colorado Nature Monument, Panama. Methods Bats were sampled over a 2‐year period on 11 islands as well as at forest‐edge and interior sites on adjacent mainland, resulting in > 8400 captures. Results The islands harboured a less diverse and structurally simplified phyllostomid bat fauna. Islands far from the mainland were especially species‐poor. This decline in species richness was associated with compositional shifts towards assemblages strongly dominated by frugivores with good dispersal abilities. Members of other ensembles, most importantly gleaning animalivores, were much less common or absent. Although overall species composition was not significantly altered, species richness at continuous forest‐edge sites was significantly lower compared with that at interior sites. Distance from the mainland and amount of forest cover in the landscape were the best predictors of species richness and assemblage composition. Responses were scale‐dependent. At the local scale, species richness was independent of island area but was correlated positively with distance from the mainland. In contrast, area effects became more important at larger spatial scales, suggesting that many species use multiple fragments. Main conclusions Our results underline the conservation value of small habitat remnants, which, even when embedded in a hostile matrix, can support a relatively diverse bat fauna, provided that there is a low degree of patch isolation and spatial proximity to larger tracts of continuous forest. Although the results at the assemblage level were inconclusive, we demonstrate that certain bat species and ensembles, particularly gleaning animalivores, exhibit high edge‐sensitivity. Our results point to habitat loss rather than changes in landscape configuration as the main process after isolation underlying phyllostomid bat responses, suggesting that conservation efforts should focus on habitat preservation instead of trying to minimize fragmentation per se at the expense of habitat amount.     

Habitat quality is more important than matrix quality for bird communities in protected areas

Protected areas are meant to preserve native local communities within their boundaries, but they are not independent from their surroundings. Impoverished habitat quality in the matrix might influence the species composition within the protected areas through biotic homogenization. The aim of this study was to determine the impacts of matrix quality on species richness and trait composition of bird communities from the Finnish reserve area network and whether the communities are being subject of biotic homogenization due to the lowered quality of the landscape matrix. We used joint species distribution modeling to study how characteristics of the Finnish forest reserves and the quality of their surrounding matrix alter species and trait compositions of forest birds. The proportion of old forest within the reserves was the main factor in explaining the bird community composition, and the bird communities within the reserves did not strongly depend on the quality of the matrix. Yet, in line with the homogenization theory, the beta‐diversity within reserves embedded in low‐quality matrix was lower than that in high‐quality matrix, and the average abundance of regionally abundant species was higher. Influence of habitat quality on bird community composition was largely explained by the species' functional traits. Most importantly, the community specialization index was low, and average body size was high in areas with low proportion of old forest. We conclude that for conserving local bird communities in northern Finnish protected forests, it is currently more important to improve or maintain habitat quality within the reserves than in the surrounding matrix. Nevertheless, we found signals of bird community homogenization, and thus, activities that decrease the quality of the matrix are a threat for bird communities.     

Birds in Anthropogenic Landscapes: The Responses of Ecological Groups to Forest Loss in the Brazilian Atlantic Forest

Habitat loss is the dominant threat to biodiversity and ecosystem functioning in terrestrial environments. In this study, we used an a priori classification of bird species based on their dependence on native forest habitats (forest-specialist and habitat generalists) and specific food resources (frugivores and insectivores) to evaluate their responses to forest cover reduction in landscapes in the Brazilian Atlantic Forest. From the patch-landscapes approach, we delimited 40 forest sites, and quantified the percentage of native forest within a 2 km radius around the center of each site (from 6 - 85%). At each site, we sampled birds using the point-count method. We used a null model, a generalized linear model and a four-parameter logistic model to evaluate the relationship between richness and abundance of the bird groups and the native forest amount. A piecewise model was then used to determine the threshold value for bird groups that showed nonlinear responses. The richness and abundance of the bird community as a whole were not affected by changes in forest cover in this region. However, a decrease in forest cover had a negative effect on diversity of forest-specialist, frugivorous and insectivorous birds, and a positive effect on generalist birds. The species richness and abundance of all ecological groups were nonlinearly related to forest reduction and showed similar threshold values, i.e., there were abrupt changes in individuals and species numbers when forest amount was less than approximately 50%. Forest sites within landscapes with forest cover that was less than 50% contained a different bird species composition than more extensively forested sites and had fewer forest-specialist species and higher beta-diversity. Our study demonstrated the pervasive effect of forest reduction on bird communities in one of the most important hotspots for bird conservation and shows that many vulnerable species require extensive forest cover to persist.     

Determinants of lepidopteran community composition and species diversity in eastern deciduous forests: roles of season, eco-region and patch size

Ecologists have long recognized that factors operating at both local and regional scales influence whether a given species occurs in an ecological community. The relative roles of variables manifested at local and regional scales on community structure, however, remain an unexplored issue for many faunas. To address this question, we compared the community composition and species diversity of forest Lepidoptera between (i) large forest tracts in historically glaciated and unglaciated regions of the eastern deciduous forest in North America, and (ii) large and small forest patches within a highly fragmented forest landscape. Specifically, we used seasonally stratified sampling to test whether regional and local differences in moth communities were related to variation in stand structure and floristic composition. At the local scale, we tested three alternative hypotheses describing the effects of patch size on moth species richness: species impoverishment, species replacement, or species supplementation. Cluster analysis revealed significant compositional differences in moth communities sampled between (i) early and late seasons, (ii) glaciated and unglaciated forest eco‐regions, and (iii) large and small forest patches. Canonical correspondence analysis suggested that floristic variation at regional scales had a greater role in determining moth community composition than local vegetation or patch‐size effects. Species richness was higher in the glaciated North Central Tillplain, and was attributable to a more diverse herbaceous feeding moth assemblage. Finally, we found evidence that both species impoverishment and species replacement processes structure the moth fauna of small woodlots; the richness of moths with larvae that feed on woody plants decreased with patch area, but herbaceous feeding species increased in diversity in smaller patches. Thus, our results suggest that local and regional differences in moth community structure are mediated by differences in host‐plant resources attributable to regional biogeographic history and local differences in patch size. Because community composition appeared to be more sensitive to environmental variation than species richness, we suggest that monitoring lepidopteran species diversity in forests will not detect significant changes in species composition due to environmental change.     

Isolation,patch size and matrix effects on bird assemblages in forest reserves

Habitat fragmentation is one of the most studied topics in ecology but our knowledge is still limited particularly concerning matrix effects on species distribution in a human-dominated landscape. We tested the ability of random sampling hypothesis, colonization–extinction dynamics and matrix-related concepts to explain the variation in species richness, total bird density and community composition in old-forest bird assemblages in two contrasting landscapes. We collected data on breeding bird abundances from 66 old-growth forest reserves in NE Finland and six larger areas in adjacent Russian Karelia using the line transect method. In Finland, protected old-forest patches are embedded in a matrix dominated by young regeneration stands. In Russia, study areas were situated in continuous, old forest dominated landscapes. Bird assemblages in old-forest patches embedded in human-modified matrix in Finland were not random samples from Russian bird assemblages. In the Finnish assemblages, species richness was lower and total bird density higher. Species richness declined with increasing distance (isolation) to Russia. Bird assemblages in large forest reserves in Finland close to Russia were structurally more similar to assemblages in the continuous reference landscape than those in small and more distant reserves. The results support the idea that several mechanisms related to colonisation–extinction dynamics and to matrix resource availability influence species distribution in fragmented landscapes but in species-specific ways. We conclude that even though small and isolated protected areas may foster high relative bird species density their ecological integrity is compromised, and therefore, improving matrix quality around reserves may lead to considerable conservation benefits.     

Effect of climate,topography and habitat on species-richness of breeding birds in Poland

We developed predictive maps of farmland and forest species richness which are based on the “focal species concept” and remote sensing data. We explored environmental preferences of 17 farmland and 33 forest species in Poland, using data from the Pan-European Monitoring Scheme. The largest number of farmland species was noted in the central and eastern parts, while the opposite trend was observed in the case of forest species, where the preferred areas were in the north and south-east. The most important environmental component affecting the bird faunas of farmland and forest was a gradient from arable fields to forest. This relationship is unimodal, which means that on a meso-geographical scale the highest species richness occurred in heterogeneous landscapes where fields are interspersed with forest. Our results also indicate that the geographical gradient in Poland's bird fauna can simply be attributed to the habitat-based distinction: western and central parts are dominated by large agricultural habitats and coniferous forest, while in other areas predominate mixed forests and extensively used farmland. Among the climatic factors, only rainfall influenced the farmland bird species. Its effect is non-linear, but positive, which means that in areas with higher rainfall more species are recorded.Our predictive maps are complementary to research on population trends, and can be an essential tool for effective management and conservation of species populations on a trans-national scale.     

Predicting distribution of <Emphasis Type="Italic">Zygaena</Emphasis> moths on West Mediterranean islands. Implications for biogeography and conservation (Lepidoptera Zygaenidae)

In this paper a conservation biogeography approach has been applied to predict distribution and conservation priorities in West Mediterranean Zygaena moths. The presence/absence matrix data for mainland region and for the three largest islands (Sardinia, Corsica and Sicily) has been analyzed. The analyses have been performed on individual species and revealed that species distribution in the Mediterranean mainland can be largely predicted. However, Sardinia and Corsica islands showed highly impoverished faunas while Sicily did not revealed a lower richness than predicted. Logistic regressions at individual species level showed that several taxa, predicted to be present in Sardinia and Corsica are actually absent. For Sicily the opposite trend was obtained and several unpredicted species actually occur on this island. Conversely to butterflies, ecological traits did not differ between predicted and unpredicted Sicilian species, suggesting that ancient and stochastic colonisations are responsible for the occurrence of several species. Due to the very low probabilities that relict populations could re-colonize islands following possible extinctions, they are suggested to deserve particular conservation efforts. In particular, Z. orana from Sardinia, Z. corsica from Corsica and Z. trifolii, Z. purpuralis and Z. carniolica from Sicily showed distinctive biogeographic patterns and/or particular rarity.     

Macro‐scale bird species richness patterns of the East Asian mainland and islands: energy,area and isolation

Aim To create a map of bird species richness (BSR) in East Asia and to examine the effect of area, isolation, primary productivity, topographic heterogeneity, and human population density on BSR. Location East Asia (from 70° E to 180° E longitude), including the eastern half of the Palaearctic Region, the entire Oriental Region, and the entire Wallacea Subregion. Methods The breeding ranges of 2406 terrestrial bird species were mapped and overlaid to create a species richness map. The BSR map was transformed into a 100 × 100 km quadrat system, and BSR was analysed in relation to land area, average normalized difference vegetation index (NDVI), elevation range, and average population density. Results In general, BSR declined from the Tropics to the Arctic. In mainland East Asia, however, BSR was highest around the Tropic of Cancer, and fluctuated between 30° and 50° N. Islands had lower BSR than adjacent mainland areas. The NDVI was strongly positively correlated with BSR in mainland areas and on islands. For mainland areas, NDVI explained 65% of the BSR variation, and topographic heterogeneity explained an additional 6% in ordinary least‐squares regression. On islands, NDVI explained 66% of BSR variation, island area explained 13%, and distance to mainland accounted for 1%. Main conclusions In East Asia, we suggest that primary productivity is the key factor underpinning patterns of BSR. Primary productivity sets the upper limits of the capacity of habitats to support bird species. In isolated areas such as islands and peninsulas, however, BSR might not reach the richness limits set by primary productivity because the degree of isolation and area size also can affect species richness. Other factors, such as spatial heterogeneity, biotic interactions, and perturbations, may also affect species richness. However, their effects are secondary and are not as strong as primary productivity, isolation, and area size.     

Living on the edge: British and Irish woodland birds in a European context

This paper reviews broad geographical patterns in the species composition of breeding woodland bird communities from Ireland to eastern Europe and outlines how processes affecting woodland birds in Britain and Ireland may differ from those operating in mainland Europe. Bird communities in British and Irish woodlands consist of a subset of the species occurring within European forests at similar latitudes. The occurrence of virtually all groups of forest birds is lower in Britain, and strikingly lower in Ireland, than in other temperate areas of mainland Europe. This phenomenon appears to form part of a west–east gradient in species diversity and is probably not just a consequence of insularity. Across this gradient there appears to be broad geographical constancy in the types (taxonomic, ecological and life-history groups) of species present. There is considerable spatial variation in habitat use by forest species within Europe. Some species in Britain probably use habitats in different ways to elsewhere for reasons related to competition, predation and historical adaptation to landscape change. Several species appear to reach the limits of their geographical ranges within Britain (i.e. in the absence of physical barriers). We suggest that range contraction of one of these species, the Common Nightingale Luscinia megarhynchos , in Britain may partially reflect redistribution into the highest quality areas in response to a wider population decline. It is argued that conclusions drawn from studies of forest birds in Britain do not necessarily apply in other regions and vice versa. There is a need for large-scale studies in Europe of the spatial variation in organization of forest bird assemblages, habitat use and the genetic structure of populations.     

Effect of forest fragmentation on bird species richness in Papua New Guinea

Tropical forests worldwide are being fragmented at a rapid rate, causing a tremendous loss of biodiversity. Determining the impacts of forest disturbance and fragmentation on tropical biotas is therefore a central goal of conservation biology. We focused on bird communities in the interior (>100 m from forest edge) of forest fragments (300, 600, and 1200 ha) in the lowlands of Papua New Guinea and compared them with those in continuous forest. We surveyed bird communities using point counts, mist‐netting, and random walks, and measured habitat and microclimate characteristics at each site. We also surveyed leaf‐dwelling arthropods, butterflies, and ants, and obtained diet samples from birds to examine food availability and food preferences. We recorded significantly fewer bird species per point in the 300‐ha forest fragment than in other study sites. Overall, we recorded 80, 84, and 88 species, respectively, in forest fragments, and 102 in continuous forest. Frugivores (especially large frugivores) and insectivores had lower species richness in forest fragments than continuous forest. Our results did not support the food scarcity hypothesis, that is, the decline of insectivorous birds in forest fragments is caused by an impoverished invertebrate prey base. We also found no significant differences among forest fragments and continuous forest in microclimates of forest interiors. Rather, we found that microhabitats preferred by sensitive birds (i.e., 30% of species with the strongest preferences for continuous forest) were less common in forest fragments (19%–31% of points) than in continuous forest (86% of points). Our results suggest that changes in microhabitats may make forest fragments unsuitable for sensitive species. However, limited dispersal capabilities could also make some species of birds less likely to disperse and occupy fragments. In addition, impoverished food resources, size of the forest fragment, or hunting pressure could contribute to the absence of large frugivorous birds in forest fragments. The forest fragments in our study, preserved as village‐based protected areas, were not large enough to sustain the bird communities found in continuous forest. However, because these fragments still contained numerous bird species, preservation of such areas can be an important component of management strategies to conserve rainforests and birds in Papua New Guinea.